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Stock-Based Compensation - Accounting for Stock-based Compensation (Details) - USD ($)
$ / shares in Units, $ in Thousands
12 Months Ended
Dec. 31, 2020
Dec. 31, 2019
Dec. 31, 2018
Share-based Compensation Arrangement by Share-based Payment Award      
Percentage of share-based compensation expense 15.00%    
Weighted average grant date fair value of options granted during the period (per share) $ 1.25 $ 1.64 $ 7.00
Stock-based compensation $ 3,641 $ 3,845 $ 5,674
Research and development      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 761 1,348 1,851
Research and development | Employee Stock Purchase Plan      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 88 36 71
Research and development | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 673 1,312 1,780
General and administrative      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 2,880 2,497 3,799
General and administrative | Employee Stock Purchase Plan      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation 9 20 48
General and administrative | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation $ 2,871 $ 2,477 3,751
Restructuring costs      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation     24
Restructuring costs | Equity Incentive Plans      
Share-based Compensation Arrangement by Share-based Payment Award      
Stock-based compensation     $ 24